Dana L. Royer

TARGET ATMOSPHERIC CO2: WHERE SHOULD HUMANITY AIM?

Paleoclimate data show that climate sensitivity is ~3 deg-C for doubled CO2, including only fast feedback processes. Equilibrium sensitivity, including slower surface albedo feedbacks, is ~6 deg-C for doubled CO2 for the range of climate states between glacial conditions and ice-free Antarctica. Decreasing CO2 was the main cause of a cooling trend that began 50 million years ago, large scale glaciation occurring when CO2 fell to 450 +/- 100 ppm, a level that will be exceeded within decades, barring prompt policy changes. If humanity wishes to preserve a planet similar to that on which civilization developed and to which life on Earth is adapted, paleoclimate evidence and ongoing climate change suggest that CO2 will need to be reduced from its current 385 ppm to at most 350 ppm. The largest uncertainty in the target arises from possible changes of non-CO2 forcings. An initial 350 ppm CO2 target may be achievable by phasing out coal use except where CO2 is captured and adopting agricultural and forestry practices that sequester carbon. If the present overshoot of this target CO2 is not brief, there is a possibility of seeding irreversible catastrophic effects.

CO 2 as a primary driver of Phanerozoic climate

Royer et al. (2004) introduce a seawater pH correction to the Phanerozoic temperature reconstruction based on δO variations in marine fossils. Although this correction is a novel idea and it is likely to have played some role in offsetting the δO record, we show that (a) The correction cannot be as large as claimed by Royer et al. (b) Irrespective of the size of the correction, a CO2 signature cannot possibly be seen in the data. (c) Even though the CO2 signature cannot be seen, the pH correction implies only a somewhat higher global temperature sensitivity than that in Shaviv and Veizer (2003), a sensitivity that is consistent with a “black body Earth”, but only marginally with the lower limit of the IPCC range.

Three keys to the radiation of angiosperms into freezing environments

Early flowering plants are thought to have been woody species restricted to warm habitats. This lineage has since radiated into almost every climate, with manifold growth forms. As angiosperms spread and climate changed, they evolved mechanisms to cope with episodic freezing. To explore the evolution of traits underpinning the ability to persist in freezing conditions, we assembled a large species-level database of growth habit (woody or herbaceous; 49,064 species), as well as leaf phenology (evergreen or deciduous), diameter of hydraulic conduits (that is, xylem vessels and tracheids) and climate occupancies (exposure to freezing). To model the evolution of species’ traits and climate occupancies, we combined these data with an unparalleled dated molecular phylogeny (32,223 species) for land plants. Here we show that woody clades successfully moved into freezing-prone environments by either possessing transport networks of small safe conduits and/or shutting down hydraulic function by dropping leaves during freezing. Herbaceous species largely avoided freezing periods by senescing cheaply constructed aboveground tissue. Growth habit has long been considered labile, but we find that growth habit was less labile than climate occupancy. Additionally, freezing environments were largely filled by lineages that had already become herbs or, when remaining woody, already had small conduits (that is, the trait evolved before the climate occupancy). By contrast, most deciduous woody lineages had an evolutionary shift to seasonally shedding their leaves only after exposure to freezing (that is, the climate occupancy evolved before the trait). For angiosperms to inhabit novel cold environments they had to gain new structural and functional trait solutions; our results suggest that many of these solutions were probably acquired before their foray into the cold.

Phanerozoic atmospheric CO2 change: evaluating geochemical and paleobiological approaches

Abstract The theory and use of geochemical modeling of the long-term carbon cycle and four paleo-PCO 2 proxies are reviewed and discussed in order to discern the best applications for each method. Geochemical models provide PCO 2 predictions for the entire Phanerozoic, but most existing models present 5–10 m.y. means, and so often do not resolve short-term excursions. Error estimates based on sensitivity analyses range from ±75–200 ppmV for the Tertiary to as much as ±3000 ppmV during the early Paleozoic. The δ 13 C of pedogenic carbonates provide the best proxy-based PCO 2 estimates for the pre-Tertiary, with error estimates ranging from ±500–1000 ppmV. Pre-Devonian estimates should be treated cautiously. Error estimates for Tertiary reconstructions via this proxy are higher than other proxies and models (±400–500 ppmV), and should not be solely relied upon. We also show the importance of measuring the δ 13 C of coexisting organic matter instead of inferring its value from marine carbonates. The δ 13 C of the organic remains of phytoplankton from sediment cores provide high temporal resolution (up to 10 3 –10 4 year), high precision (±25–100 ppmV for the Tertiary to ±150–200 ppmV for the Cretaceous) PCO 2 estimates that can be near continuous for most of the Tertiary. Its high temporal resolution and availability of continuous sequences is advantageous for studies aiming to discern short-term excursions. This method, however, must correct for changes in growth rate and oxygen level. At elevated PCO 2 (∼750–1250 ppmV), this proxy loses its sensitivity and is not useful. The stomatal density and stomatal index of land plants also provide high temporal resolution ( 2 year), high precision (±10–40 ppmV for the Tertiary and possibly Cretaceous) PCO 2 estimates, and so also is ideal for discerning short-term excursions. Unfortunately, this proxy also loses sensitivity at some level of PCO 2 above 350 ppmV (which, currently, is largely undetermined). Our analysis of the recently developed δ 11 B technique shows that it currently is not yet well constrained. Most importantly, it requires the assumption that the boron isotopic composition of the ocean remains nearly constant through time. In addition, it assumes that there are no biological or temperature effects and that diagenetic alteration of the boron isotopic composition does not occur. A fifth CO 2 proxy, based on the redox chemistry of marine cerium, has several fundamental flaws and is not discussed in detail here.

Sensitivity of leaf size and shape to climate: global patterns and paleoclimatic applications

• Paleobotanists have long used models based on leaf size and shape to reconstruct paleoclimate. However, most models incorporate a single variable or use traits that are not physiologically or functionally linked to climate, limiting their predictive power. Further, they often underestimate paleotemperature relative to other proxies. • Here we quantify leaf-climate correlations from 92 globally distributed, climatically diverse sites, and explore potential confounding factors. Multiple linear regression models for mean annual temperature (MAT) and mean annual precipitation (MAP) are developed and applied to nine well-studied fossil floras. • We find that leaves in cold climates typically have larger, more numerous teeth, and are more highly dissected. Leaf habit (deciduous vs evergreen), local water availability, and phylogenetic history all affect these relationships. Leaves in wet climates are larger and have fewer, smaller teeth. Our multivariate MAT and MAP models offer moderate improvements in precision over univariate approaches (± 4.0 vs 4.8°C for MAT) and strong improvements in accuracy. For example, our provisional MAT estimates for most North American fossil floras are considerably warmer and in better agreement with independent paleoclimate evidence. • Our study demonstrates that the inclusion of additional leaf traits that are functionally linked to climate improves paleoclimate reconstructions. This work also illustrates the need for better understanding of the impact of phylogeny and leaf habit on leaf-climate relationships.

The evolution and functional significance of leaf shape in the angiosperms

Angiosperm leaves manifest a remarkable diversity of shapes that range from developmental sequences within a shoot and within crown response to microenvironment to variation among species within and between communities and among orders or families. It is generally assumed that because photosynthetic leaves are critical to plant growth and survival, variation in their shape reflects natural selection operating on function. Several non-mutually exclusive theories have been proposed to explain leaf shape diversity. These include: thermoregulation of leaves especially in arid and hot environments, hydraulic constraints, patterns of leaf expansion in deciduous species, biomechanical constraints, adaptations to avoid herbivory, adaptations to optimise light interception and even that leaf shape variation is a response to selection on flower form. However, the relative importance, or likelihood, of each of these factors is unclear. Here we review the evolutionary context of leaf shape diversification, discuss the proximal mechanisms that generate the diversity in extant systems, and consider the evidence for each the above hypotheses in the context of the functional significance of leaf shape. The synthesis of these broad ranging areas helps to identify points of conceptual convergence for ongoing discussion and integrated directions for future research.

Climate sensitivity constrained by CO2 concentrations over the past 420 million years.

A firm understanding of the relationship between atmospheric carbon dioxide concentration and temperature is critical for interpreting past climate change and for predicting future climate change. A recent synthesis suggests that the increase in global-mean surface temperature in response to a doubling of the atmospheric carbon dioxide concentration, termed ‘climate sensitivity’, is between 1.5 and 6.2 °C (5–95 per cent likelihood range), but some evidence is inconsistent with this range. Moreover, most estimates of climate sensitivity are based on records of climate change over the past few decades to thousands of years, when carbon dioxide concentrations and global temperatures were similar to or lower than today, so such calculations tend to underestimate the magnitude of large climate-change events and may not be applicable to climate change under warmer conditions in the future. Here we estimate long-term equilibrium climate sensitivity by modelling carbon dioxide concentrations over the past 420 million years and comparing our calculations with a proxy record. Our estimates are broadly consistent with estimates based on short-term climate records, and indicate that a weak radiative forcing by carbon dioxide is highly unlikely on multi-million-year timescales. We conclude that a climate sensitivity greater than 1.5 °C has probably been a robust feature of the Earth’s climate system over the past 420 million years, regardless of temporal scaling.

Correlations of climate and plant ecology to leaf size and shape: potential proxies for the fossil record

The sizes and shapes (physiognomy) of fossil leaves are widely applied as proxies for paleoclimatic and paleoecological variables. However, significant improvements to leaf-margin analysis, used for nearly a century to reconstruct mean annual temperature (MAT), have been elusive; also, relationships between physiognomy and many leaf ecological variables have not been quantified. Using the recently developed technique of digital leaf physiognomy, correlations of leaf physiognomy to MAT, leaf mass per area, and nitrogen content are quantified for a set of test sites from North and Central America. Many physiognomic variables correlate significantly with MAT, indicating a coordinated, convergent evolutionary response of fewer teeth, smaller tooth area, and lower degree of blade dissection in warmer environments. In addition, tooth area correlates negatively with leaf mass per area and positively with nitrogen content. Multiple linear regressions based on a subset of variables produce more accurate MAT estimates than leaf-margin analysis (standard errors of ±2 vs. ±3°C); improvements are greatest at sites with shallow water tables that are analogous to many fossil sites. The multivariate regressions remain robust even when based on one leaf per species, and the model most applicable to fossils shows no more signal degradation from leaf fragmentation than leaf-margin analysis.

Depth to pedogenic carbonate horizon as a paleoprecipitation indicator

An analysis of 1168 modern soil profiles from the U.S. Natural Resources Conservation Service yields no correlation between mean annual precipitation and depth to the top of the carbonate horizon ( r 2 = 0.03; p < 0.001). Parent material and soil texture both play negligible roles in this regression. When combined with similar published studies (n = 1481), r 2 improves slightly ( r 2 = 0.31; p < 0.001). Caution is therefore advised in using this or any previously published regression for inferring paleoprecipitation from paleosols. However, in the combined data set, carbonate horizon‐bearing soils correlate with a mean annual precipitation of < 760 mm ( p < 0.05). Thus the presence vs. absence of pedogenic calcium carbonate is a good indicator of paleoprecipitation above or below this value.

Sharply increased insect herbivory during the Paleocene–Eocene Thermal Maximum

The Paleocene–Eocene Thermal Maximum (PETM, 55.8 Ma), an abrupt global warming event linked to a transient increase in pCO2, was comparable in rate and magnitude to modern anthropogenic climate change. Here we use plant fossils from the Bighorn Basin of Wyoming to document the combined effects of temperature and pCO2 on insect herbivory. We examined 5,062 fossil leaves from five sites positioned before, during, and after the PETM (59–55.2 Ma). The amount and diversity of insect damage on angiosperm leaves, as well as the relative abundance of specialized damage, correlate with rising and falling temperature. All reach distinct maxima during the PETM, and every PETM plant species is extensively damaged and colonized by specialized herbivores. Our study suggests that increased insect herbivory is likely to be a net long-term effect of anthropogenic pCO2 increase and warming temperatures.